Multiple mechanisms sustain a plant-animal facilitation on a coastal ecotone

Theory suggests that species distributions are expanded by positive species interactions, but the importance of facilitation in expanding species distributions at physiological range limits has not been widely recognized. We investigated the effects of the nurse shrub Tamarix chinensis on the crab Helice tientsinensis on the terrestrial borders of salt marshes, a typical coastal ecotone, where Tamarix and Helice were on their lower and upper elevational distribution edges, respectively. Crab burrows were abundant under Tamarix, but were absent in open areas between Tamarix. Removing Tamarix decreased associated crab burrows with time, while simulating Tamarix in open areas by shading, excluding predators, and adding Tamarix branches as crab food, increased crab burrows. Measurements of soil and microclimate factors showed that removing Tamarix increased abiotic stress, while simulating Tamarix by shading decreased abiotic stress. Survival of tethered crabs was high only when protected from desiccation and predation. Thus, by alleviating abiotic and biotic stresses, as well as by food provision, Tamarix expanded the upper intertidal distribution of Helice. Our study provides clear evidence for the importance of facilitation in expanding species distributions at their range limits, and suggests that facilitation is a crucial biological force maintaining the ecotones between ecosystems.